Biodiversity studies in Afrotropical moth flies (Diptera: Psychodidae)

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BIODIVERSITY STUDIES IN AFROTROPICAL MOTH FLIES

(DIPTERA: PSYCHODIDAE)

Gunnar Mikalsen Kvifte

Master thesis in biology - Biodiversity, Evolution and Ecology

University of Bergen 2011

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I found it and I named it, being versed in taxonomic latin, thus became Godfather of an insect, and its first describer, and I want no other fame.

-Vladimir Nabokov

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First of all I would like to thank my supervisors Trond Andersen and Richard Telford for their excellent guidance and encouragement through the thesis process. Trond was the first to suggest Psychodidae as a worthy subject for study, and has remained extremely helpful in writing, labwork and illustration throughout my education. Richard helped me planning fieldwork and has shared his considerable methodological expertise generously. I can not thank them enough for their contributions to this project.

I am very grateful to everybody involved in the MATRIX project for support during the fieldwork in Uganda. Without all the advice and technical help I received from Vigdis Vandvik, Josephine Esaete and Brooke Wilkerson I would have had a considerably harder time in Uganda. I also could not have done without my field assistant Afeku Alfred, whose wide ecological and botanical knowledge of Budongo forest was crucial in my site selection. For further help, advice and company during my stay at the Budongo Conservation Field Station I thank Roman M. Wittig, Zephyr Kiwede, Tonny Kidega and all the students and staff of the BCFS. Asanti sana!

The staff at the natural history museums in Makerere: Robert Kityo, Perpetra Akite and Fabiano Ebonga and Bergen: Per Djursvoll, Katrine Kongshavn, Gladys Ramirez and Steffen Roth were very helpful in solving technical and/or administrative problems. I also want to thank Marc de Meyer and Eliane de Coninck of the Royal Museum for Central Africa, Tervuren for the loan of the Trichomyia congoensis holotype, and Erica McAlister at the British Museum of Natural History, London for guidance during my visit in the entomology collections in London.

Discussions with Gregory R. Curler were of great help in resolving some difficult taxonomical issues; I am especially grateful for his comments on an earlier draft of chapter 2. I also appreciated good advice from Torbjørn Ekrem, Jan Ježek, Bjarte Jordal, Lawrence Kirkendall, Humberto F.

Mendes, Torstein Solhøy, Gaute Velle and Endre Willassen. Furthermore, I want to thank Louis Boumans, Freddy Bravo, Cínthia Chagas, Øyvind Håland, Jan Ježek and Salih Krek for providing me with copies of some of the more obscure taxonomic publications.

Less academic kinds of support from my friends and family has been of great help – writing a thesis is a stressful and largely lonely process which I would have great difficulties with if it was not for you. Svein, Lasse, Inge, the entire «fagpils» community, my fellow Klubb Fantoft-ers – you are all greatly appreciated. See you at Baran for the thesis celebration!

Finally I have to thank my lovely Aleksandra for her everlasting patience and loving support, which has been and remains very important to me.

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We are living in an age of extinction. Currently, species are disappearing at rates that are believed to be hundreds or even thousands of times higher than the «natural» rate of extinction; principally because of habitat loss induced by changes in land use and climate (Pimm et al. 1995; Sala et al. 2000). A majority of the extinctions are likely to go unnoticed as most of the endangered biodiversity consists of unknown or poorly known tropical insect species (Lewis & Basset 2007).

Our ignorance of tropical insect diversity is immense - up to 95% of the multicellular species on Earth may remain unknown, and for most of the described species the only available information is basic data about the morphology and geographical origin of a few specimens (Stork 2007). Even less is known about species assemblages and biological communities and of the processes that shape them (Godfray et al. 1999).

Accompanying the alarming rates of species extinction is a severe loss of taxonomic knowledge and know-how; the so-called taxonomic impediment (Hoagland 1996). The number of active taxonomic specialists has decreased steadily the last 50 years, both among amateurs and professionals (Hopkins & Freckleton 2002; Wheeler 2004). Within the same time frame, however, the relevance of taxonomy has, if anything, increased! Firstly, our ignorance has been proven to be even more extensive than previously thought. This has been revealed both by new sampling methods (e.g. canopy fogging; Erwin 1982; Thunes et al. 2004) and methodological advances in species identification (e.g. use of DNA to resolve cryptic species complexes: Hebert et al. 2004; Smith et al. 2006). Just as notably, taxonomy provides an important basis for most other subdisciplines of biology, including but not limited to conservation biology (e.g. Dubois 2003; Mace 2004), forestry science (e.g. Eidt 1995), agricultural sciences (e.g. Rosen 1986), palaeoecology (e.g. Birks 1994), disease vector control (e.g. Brooks & Hoberg 2001; Van Bortel et al. 2001) and biomonitoring (e.g.

Terlizzi et al. 2003).

In this thesis I am attacking one of the biggest unsolved science questions of our time – the magnitude and formation of tropical insect biodiversity. Following the recommendations of Godfray et al. (1999) and Gotelli (2004), I will do this using a combination of revisionary alpha taxonomy and statistical interpolation from species inventories. My focus group is the Psychodidae, a species-rich family of small, hairy flies with a near-cosmopolitan distribution.

In chapter 1, I provide a systematic context for the work; reviewing the diversity, morphology, systematics and classification of Psychodidae of the world. Six subfamilies Bruchomyiinae, Phlebotominae, Psychodinae, Sycoracinae, Trichomyiinae and Horaiellinae are recognised. The chapter gives an overview of their characteristics, species diversity and known biology. The chapter also includes a broad review of the morphological characters used in the taxonomy of Psychodidae with an emphasis on the most species-rich subfamily Psychodinae.

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Chapter 2 catalogues the Afrotropical fauna of non-phlebotomine Psychodidae. A total of 175 species in 27 genera are listed, all with full bibliographic citations and distributional data.

Cryptotelmatoscopus Vaillant, 1982 is placed as a subgenus of Clogmia Enderlein, 1935, stat.

nov.; Karakovounimerus Ježek, 1990 is placed as a subgenus of Panimerus Eaton, 1904, stat.

nov.; Orgaoclogmia Ježek & van Harten, 1996 is synonymised with Cryptotelmatoscopus Vaillant, 1982, syn.nov.; and Rhipidopsychoda Vaillant, 1991 is synonymised with Threticus Eaton, 1904 syn.nov. Telmatoscopus flagellifer Freeman, 1949, Mormia soelii Wagner & Andersen, 2007 and Rhadinoscopus triangulatus Wagner, 1979 are transferred to Hemimormia Krek, 1971, comb.nov.;

Orgaoclogmia caboverdeana Ježek & van Harten, 1996 is transferred to Clogmia Enderlein, 1937, comb.nov.; Telmatoscopus pilosternatus Satchell, 1955 is transferred to Mormopericomiella Ježek

& van Harten, 2002, comb.nov.; Copropsychoda bulbosa Ježek & van Harten, 2005, Falsologima verrucosa Ježek & van Harten, 2005, Psychana rujumensis Ježek & van Harten, 2005 and Psychodocha khoralkhwairensis Ježek & van Harten, 2009 are transferred to Psychoda Latreille, 1796 comb.nov.;

and Psychoda boettgeri Wagner, 1979 is transferred to Threticus Eaton, 1904 comb.nov.

Chapter 3 investigates the habitats of Psychodidae in Budongo forest, Uganda, based on Malaise trap samples in the forest’s four different vegetation types: Colonizing forest, mixed mature forest, Cynometra alexandrei-dominated climax forest and swamp forest. A total of 546 specimens were collected, of which 103 specimens of 38 morphospecies in eight genera could be identified.

One of the genera appears to be new to science and five other genera are recorded for the first time from Uganda. The genus Neotelmatoscopus Tonnoir, 1933, is recorded for the first time from the Afrotropical region. The dominant genus both in species richness and in total abundance is Psychoda. The colonizing and mixed mature forest types had quite similar species assemblages and an apparently much higher diversity than both the swamp forest and the Cynometra forest. The results are briefly discussed and compared to a similar study in Brazil.

Chapter 4 is a review of the Afrotropical species of Trichomyia Haliday in Curtis, 1839.

Trichomyia piricornis Freeman, 1949 and Trichomyia congoensis Satchell, 1956 are redescribed based on type material. Trichomyia nodosa Duckhouse, 1980, Trichomyia dlinzae Duckhouse, 1980 and Trichomyia brochata Quate, 1957 are diagnosed. Trichomyia anderseni, sp.nov., Trichomyia budongoensis sp.nov., Trichomyia cornifera sp.nov., Trichomyia cynometrae sp.nov. and Trichomyia telfordi sp.nov. are described as new to science. A key to the males of Afrotropical Trichomyia is provided and the genus’ biogeography and phylogeny is briefly discussed.

The results presented in this thesis are only a small step towards a more thorough understanding of Afrotropical Psychodidae diversity. As indicated in chapter 2, many taxa are in need of revision;

notably Threticus, Telmatoscopus s.l., Psychoda and Mormiina. Chapter 3 makes it clear that taxonomy of female Psychodidae should be given priority – both exploring their morphology more closely both for genus- and species-level characters and associating females and males using e.g.

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show a relatively clear biogeographic pattern, which can be tested through further sampling and phylogenetic studies of African and other Trichomyia.

Ecological questions worth pursuing further include the specific natural history of the immature stages of individual species, which would facilitate the search for autecological patterns in diversity. Furthermore, species inventories should be made along comparable land use gradients in other tropical sites, both in the Afrotropics and in other regions. Comparing and contrasting Psychodids with other semiaquatic decomposer taxa (e.g. Chironomidae: Orthocladiinae) could be interesting to see whether the observed patterns given in chapter 3 can be linked to feeding guilds.

DISCLAIMER

In line with the International Code of Zoological Nomenclature (ICZN) articles 8.2 and 8.3, this thesis is not published for nomenclatural purposes. The nomenclatural acts in this thesis are therefore not yet valid by ICZN criteria and should be considered mere suggestions rather than actual scientific names. The taxonomic decisions herein will be validated in later publications satisfying the criteria in ICZN art. 8.

REFERENCES

Birks, H. J. B. (1994) The importance of pollen and diatom taxonomic precision in quantitative palaeoenvironmental reconstructions. Review of Palaeobotany and Palynology, 83, 107- 117.

Brooks, D. R. & Hoberg, E. P. (2001) Parasite systematics in the 21st century: opportunities and obstacles. Trends in Parasitology, 17, 273-275.

Dubois, A. (2003) The relationships between taxonomy and conservation biology in the century of extinctions. Comptes Rendus Biologies, 326, supplement 1, 9-21.

Eidt, D. C. (1995) The Importance of Insect Taxonomy and Biosystematics to Forestry. Forestry Chronicle, 71, 581-583.

Erwin, T. L. (1982) Tropical Forests: Their Richness in Coleoptera and Other Arthropod Species.

The Coleopterists Bulletin, 36, 74-75.

Godfray, H. C. J., Lewis, O. T. & Memmott, J. (1999) Studying insect diversity in the tropics.

Philosophical Transactions of the Royal Society of London (B), 354, 1811-1824.

Gotelli, N. J. (2004) A taxonomic wish-list for community ecology. Philosophical Transactions of the Royal Society of London (B), 359, 585-597.

Hebert, P. D. N., Penton, E. H., Burns, J. M., Janzen, D. & Hallwachs, W. (2004) Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences of the United States of America,

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101, 14812-14817.

Hoagland, K. E. (1996) The taxonomic impediment and the Convention of Biodiversity. Association of Systematics Collections Newsletter, 24, 61-62, 66-67.

Hopkins, G. W. & Freckleton, R. P. (2002) Declines in the numbers of amateur and professional taxonomists: implications for conservation. Animal Conservation, 5, 245-249.

International Commission for Zoological Nomenclature (1999). International Code of Zoological Nomenclature. 4^th edition. Available online from http://www.nhm.ac.uk/hosted-sites/iczn/code/, assessed august 16^th 2011

Lewis, O. T. & Basset, Y. (2007) Insect Conservation in Tropical Forests. In: A. J. A. Stewart, T.

R. New & O. T. Lewis (Eds), Insect Conservation Biology. Royal Entomological Society of London, London, pp. 34-56.

Mace, G. M. (2004) The role of taxonomy in species conservation. Philosophical Transactions of the Royal Society of London (B), 359, 711-719.

Pimm, S. L., Russell, G. J., Gittleman, J. L. & Brooks, T. M. (1995) The Future of Biodiversity.

Science, 269, 347-350.

Rosen, D. (1986) The Role of Taxonomy in Effective Biological-Control Programs. Agriculture Ecosystems & Environment, 15, 121-129.

Sala, O. E., Chapin, F. S., Armesto, J. J., Berlow, E., Bloomfield, J., Dirzo, R., et al. (2000) Global biodiversity scenarios for the year 2100. Science, 287, 1770-1774.

Smith, M. A., Woodley, N. E., Janzen, D. H., Hallwachs, W. & Hebert, P. D. N. (2006) DNA barcodes reveal cryptic host-specificity within the presumed polyphagous members of a genus of parasitoid flies (Diptera: Tachinidae). Proceedings of the National Academy of Sciences of the United States of America, 103, 3657-3662.

Stork, N. E. (2007) Biodiversity - World of insects. Nature, 448, 657-658.

Terlizzi, A., Bevilacqua, S., Fraschetti, S. & Boero, F. (2003) Taxonomic sufficiency and the increasing insufficiency of taxonomic expertise. Marine Pollution Bulletin, 46, 556-561.

Thunes, K. H., Skartveit, J., Gjerde, I., Starý, J., Solhøy, T., Fjellberg, A., et al. (2004) The arthropod community of Scots pine (Pinus sylvestris L.) canopies in Norway. Entomologica Fennica, 15, 65-90.

Van Bortel, W., Harbach, R. E., Trung, H. D., Roelants, P., Backeljau, T. & Coosemans, M. (2001) Confirmation of Anopheles varuna in Vietnam, previously misidentified and mistargeted as the malaria vector Anopheles minimus. American Journal of Tropical Medicine and Hygiene, 65, 729-732.

Wheeler, Q. D. (2004) Taxonomic triage and the poverty of phylogeny. Proceedings of the Royal Society of London (B), 359, 571-583.

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Introduction to Psychodidae

The moth flies (Diptera: Psychodidae) are a species-rich family of small, fuzzy gnats found all over the world. Most known species are considered to be aquatic or semiaquatic, but the family exhibits a large and still poorly explored ecological diversity. About 2900 species are described this far, but as the fauna in most parts of the world remains poorly studied the actual number of species is clearly much higher (see Curler & Courtney 2009).

The only area that has been investigated thoroughly for moth flies is the West Palearctic, but even here new species are still discovered at a fairly regular rate (e.g. Withers 2003; Salmela &

Piirainen 2005; Wagner & Schrankel 2005; Ježek 2006; Ježek & Hájek 2007; Beran et al. 2010).

More research is clearly needed to improve our knowledge of Psychodid diversity. Wagner &

Ibañez-Bernal (2009) estimate that the total world fauna may comprise as many as 20 000 species.

Most authors recognize the Psychodidae as a single family comprising six extant subfamilies:

Bruchomyiinae, Phlebotominae, Psychodinae, Sycoracinae, Horaiellinae and Trichomyiinae.

Some authors, however, treat the group as a superfamily Psychodoidea; comprising two families (e.g. Williams 1993, Azar et al. 1999). Under this scheme Phlebotomidae consists of the two subfamilies Bruchomyiinae and Phlebotominae while Psychodidae s.str. comprises the subfamilies Psychodinae, Horaiellinae, Sycoracinae and Trichomyiinae. As both views seem phylogenetically sound, choosing between them is a matter of personal taste.

Synapomorphies indicating that Psychodidae form a monophyletic group include dense vestiture of the body; antennae with membranous sensory filaments (ascoids); wing with reduced anal area; second basal cell and A₂ vein of wing shortened; distal part of wing without crossveins;

reduction from 3 to 2 spermathecae in the female; and male genitalia inverted 180° (Hennig 1972;

Quate & Vockeroth 1981). The sister group of the Psychodidae is less clear. Based on characters of the larval mouthparts, Wood & Borkent (1989) suggested that the moth flies’ closest relatives are a clade comprising the families Trichoceridae, Anisopodidae, Scatopsidae, Synneuridae and Perissomatidae. Oosterbroek & Courtney (1995) analysed an expanded morphological data set and resolved Psychodidae as the sister grup to a clade consisting of Tipulomorpha, Anisopodidae and

Chapter 1

Introduction to the Psychodidae

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Brachycera. Molecular data suggest a close relationship with Tanyderidae (Bertone et al. 2008); a relationship which has long been suspected but without any clear morphological synapomorphies (e.g. Hennig 1972, 1973 cited in Oosterbroek & Courtney 1995).

Because of the morphological and structural diversity within the Psychodidae, each of the six subfamilies are presented separately below.

MORPHOLOGY

The present account is meant as an introduction to the general morphology of psychodid flies, with an emphasis on the Psychodinae. Only adult stages are treated here; for information on the larvae see e.g. Tonnoir (1933), Keilin & Tate (1937), Satchell (1947, 1949, 1953), Vaillant (e.g. 1959, 1963, 1971), Duckhouse (1985, 1994), Mahmood & Alexander (1992) and Leite & Williams (1996). Some aspects of the pupae are treated by e.g. Vaillant (1971), Satchell (1948), Leite et al.

(1991) and Curler & Courtney (2009).

Morphological terminology varies slightly between different authors. The present work follows Curler & Courtney (2009).

Head (Figs 1.1-1.3)

The taxonomically most important parts of the head are the eyes, the antennae and the mouthparts.

The shape and chaetotaxy of the frons (e.g. the frontal scar patch; the arrangement of hair scars on the frons), the vertex and the clypeus are occasionally also of some value. In some groups, there are also some accessory organs (e.g. corniculi) that are taxonomically useful.

In most Psychodinae the eyes are extended between frons and vertex, creating an eyebridge (Fig. 1.1). The width and extent of this bridge is often an important taxonomic character. Other subfamilies lack this eyebridge completely (e.g. Fig. 4.4).

The sclerotized inner parts surrounding the pharynx, especially the cibarium, have been proven useful as a systematic character especially in the Phlebotominae, but has been less used in other subfamilies (Quate 1962).

In some groups the terminal lobes of the labium, called the labellum, are of importance.

In most Psychodidae these are fleshy and distinctly bulbous. However, in Phlebotominae and Psychodinae: Psychodini they are modified (Quate 1959). The appearance of the labrum and the maxillae seem rather uniform throughout the family. The maxillary palps primitively consist of five segments called palpomeres, but in most subfamilies this number is reduced to four (in Psychodinae, Sycoracinae, some Trichomyiinae) or three segments (Horaiellinae, some Trichomyiinae).

Functional mandibles are only present in Phlebotominae, Sycoracinae and possibly Horaiellinae.

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Fig. 1.1-1.3. Morphology of Psychodidae I. (1.1) Psychoda terskolina Vaillant & Joost, 1983:

Head; (1.2) Pericoma nielseni Kvifte, 2010: Flagellomeres 10-14 (fusiform); (1.3) Telmatoscopus sp.n.1, Uganda: Flagellomeres 11-14 (nodiform). ab – antennal basis, asc – ascoid, cl – clypeus, ebr – eyebridge, f – frons, in – internode, lbl – labellum, lbr – labrum, n – node, p1 – first palpomere, vrt – verticil, vx – vertex

The antennae of the Psychodidae are divided into three parts: The scape, the pedicel and a flagellum consisting of between 8 and 111 flagellomeres. The vast majority of species have 12-14 flagellomeres. The first flagellomere is occasionally called the postpedicel, especially in groups where this segment has a different shape than the other flagellomeres. The distalmost flagellomeres sometimes carry tubercles, spines or other structures of taxonomic value.

A feature unique to the Psychodidae is the presence of ascoids – hyaline sensory organs carried on the flagellomeres. The most simple ascoids are digitiform or filamentose, but a large range of variation exists for this character with modifications in branching and/or flattening being the most common changes. Additional types of sensillae are also common in many genera, most conspicuosly the so-called «bullseye organs» in some Mormiini (Vaillant 1974, p. 132) and the postpedicel spines of many Pericomaini. Less conspicuous sensillae have also been found; in a recent study of the antennae of Psychoda s.l., Faucheux & Gibernau (2011) detected no less than seven types of antennal sensillae.

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Thorax (Fig. 1.5)

Few systematic characters are found in the thorax. Some characteristics common for the group include a straight transverse suture in the scutum and the metanotum protruding into the abdomen.

The pteropleurites show some variation especially within the Psychodinae and have been suggested as an important character in tribal classification of this subfamily (Ježek 1984).

Some groups within the Psychodinae have developed more or less elaborate organs on the meso- and metanotum. They are most properly called allurement organs, although many authors use the Lepidopteran morphological terms patagia and tegulae instead (Duckhouse 1990). For most taxa the function of these organs is not clear. However, in the European species Ulomyia fuliginosa (Meigen, 1804) the anterior mesothoracal allurement organ is involved in secretion of pheromones (Elger 1981).

Fig. 1.4-1.6. Morphology of Psychodidae II. (1.4) Threticus sp.n.1, Uganda: Wing; (1.5) Telmatoscopus sp.n.1, Uganda: Thorax; (1.6) Sycorax sp. near malayensis Quate,

Malaysia: Wing. asp – anterior spiracle, cn – costal node, CuA – cubital / anal veins, cx – coxa, h – haltere, jug – jugum, M – medial veins, R – radial veins, Sc – subcosta, sct – scutum, scu – scutellum.

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Wing (Figs 1.4, 1.6)

The ground-plan wing has 10 veins reaching the margin. Five of these are branches of the radial vein (R), three are branches of the medial vein (M) and two are of the cubital vein (CuA). In the subfamilies Sycoracinae, Trichomyiinae and Horaiellinae, the R vein has only four branches (e.g.

Fig. 1.6). Also, there is a costal vein (C) surrounding the wing and a reduced subcostal vein (Sc) at the wing basis. The subfamily Psychodinae have a well-developed triangular lobe at the base of the wing; this is occasionally misnamed an alula, but is properly called the neala or jugum.

The wing forks are composed of veins R₂+R₃ (the R-fork) and M₁+M₂ (the M-fork). They are present in all Psychodidae wings, although they sometimes are incomplete at their bases. Their relative placement to each other and to CuA₂ are in much use as taxonomic characters.

Usually, Psychodid wings and bodies are covered in hair-like scales. Some groups have hairs on the wing membrane, but most moth flies have hairs confined to the wing veins only. The colour patterns formed by these hairs are presumably species-specific, but as the hairs tend to break loose in alcohol preserved material they are not much used as taxonomic characters.

Abdomen and terminalia (Fig. 1.7)

The abdomen consists of 9 sternites and 10 tergites, abbreviated S1-9 and T1-10. The terminal part including the genitalia is inverted 180 degrees in all taxa except the Sycoracine genera Sycorax Haliday in Curtis, 1839 and Parasycorax Duckhouse, 1972.

The male genitalia (Fig. 1.7) carry many of the most important systematic characters within the group, both on species level and in higher classification. They consist of the sternite 9 and tergites 9 and 10 with their appendages; which for all groups include the gonopods. Sternite 9 is usually present as a narrow band above the gonopods, called the hypandrium.

Primitively, the Psychodidae have a gonopod with two segments: a gonocoxite and a gonostylus. A few taxa have one or two additional appendages (see e.g. Duckhouse 1978 and chapter 4, this volume). The gonocoxite is articulated posteriorly to the gonostylus and is often expanded basally. On the inner surface of the gonocoxite, its expansions are called the anterior and posterior gonocoxal apodeme, respectively (Quate & Brown 2004). In this volume I have adopted the term outer gonocoxal apodeme for the ventral (morphologically dorsal) anterior gonocoxal expansions seen in many taxa. The gonocoxal apodeme of Quate & Brown (2004) is thus labelled the inner gonocoxal apodeme.

The copulatory organ of the male – the aedeagus – is divided into a basiphallus (occasionally called an aedeagal apodeme, ejaculatory apodeme or phallapodeme), a distiphallus and a subgenital plate. The distiphallus primitively consists of a symmetrical pair of phallomeres which often are called distiphalli. In many taxa these are fused or secondarily asymmetrical. The phallomeres are often flanked by paired gonapophyses and paired or unpaired parameres. In some groups, the

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parameres form a phallic sheath around the distiphalli. Additional parts of the aedeagus include the furca of many Psychodinae: Paramormiini, which articulates the aedeagus to the ventral epandrial plate (Duckhouse 1987).

Tergite 9 is called the epandrium and carries on its dorsal surface a ventral epandrial plate (ventral prior to genital rotation; Duckhouse (1987)). Posteriorly the ventral epandrial plate articulates with the 10^th tergite and the cerci. In Psychodinae, the cerci are modified to form cercopods, carrying from 1 to over 100 retinaculae usually at their apex. The shape and number of retinaculae is taxonomically important, although intraspecific variation in number is not uncommon in many groups.

The female genitalia of Psychodidae are insufficiently explored for most taxa, with the Fig. 1.7. Morphology of Psychodidae III. (1.7) Threticus sp.n.1, Uganda: Male genitalia. bas –

basiphallus, crc – cercopod, dst – distiphalli, ep – epandrium, gcx – gonocoxite, gst – gonostylus, hyp – hypandrium, iga – inner gonocoxal apodeme, pr – paramere, ret – retinaculum, t10 – The 10th tergite, vep – ventral epandrial plate

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exception of the Phlebotominae (Curler & Courtney 2009). Vaillant (1971) redescribes the female Clogmia albipunctata (Williston, 1893) in considerable detail, and Sæther (1977) illustrates two species of Psychoda Latreille, 1796. Still, a review of female genital morphology would be desireable in most Psychodid groups.

CLASSIFICATION

Bruchomyiinae Alexander, 1920

The Bruchomyiinae are often considered to be the most primitive group of Psychodidae, based on plesiomorphies in the wing venation, antennae, mouthparts and lack of eyebridges (Quate &

Alexander 2000). The group comprises 41 extant species in tropical and subtropical parts of the world, with most species known from the Neotropical Region (Williams 2003; Wagner 2006;

Santos et al. 2009b). Three genera Bruchomyia Alexander, 1920, Nemapalpus Macquart, 1838 and Eutonnoiriella Alexander, 1940 are recognized. However, it is likely that at least Nemapalpus as presently defined is an artificial assemblage (Quate & Alexander 2000).

Rarely collected, adult Bruchomyiines seem to be mostly restricted to moist forests.

Nemapalpus ledgeri Stuckenberg, 1978 and N. davidsoni Stuckenberg, 1978 were, however collected in the nests of rock hyraxes, Procavia capensis (Pallas, 1766), in an arid savanna landscape (Stuckenberg 1978). Immature stages of two species have been described, but their biology remains poorly known (Mahmood & Alexander 1992). A Nemapalpus species in New Zealand was found to develop in rotting wood (Duckhouse 1980), and Williams (2003) mentions larvae and pupae in association with ant colonies.

Synapomorphic characters for the Bruchomyiinae include the reduction from two to one spermatheca in the female and from two to one genital opening in the male, as well as non-functional mouthparts in both sexes (Hennig 1972; Wagner 2006). Other characteristic features include the long legs and slender body; maxillary palp with five segments; Sc connected to R₁ and often C;

CuA₁, CuA₂ and CuA₃ elongated and gonostyles without thorns (Duckhouse 1965; Wagner 2006).

Another possible apomorphy for the clade is a tendency to duplicate antennal segments – whereas Nemapalpus has 16 flagellomeres, Bruchomyia has around 30 and Eutonnoiriella as many as 111 (Williams 2003)! The subfamily has been suspected to be paraphyletic (Fairchild 1955; cited in Hennig 1972), but is now usually regarded as the monophyletic sister group of the Phlebotominae (Hennig 1972; Azar et al. 1999; Curler 2009).

Undisputed fossil Bruchomyiinae are from Baltic and Dominican amber; both of which are of Tertiary age (Wagner 2006). In addition, the Jurassic wing fossil Dacochile microsoma Poinar &

Brown, 2004 has been suggested as a possible member of the group (Woodley 2005).

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Phlebotominae Rondani, 1840

More study has been devoted to the Phlebotominae than to all the other Psychodid subfamilies combined. This is due to their medical importance; around 70 species in the genera Lutzomyia França, 1924 and Phlebotomus Rondani & Berte in Rondani, 1840 are important vectors of pathogenic protozoans such as Leishmania (Williams 1993; Azar & Nel 2003). Until recently, the Phlebotominae was considered to consist of six extant genera (e.g. Williams 1993). The group has, however, been the focus of several large-scale phylogenetic studies (e.g. Galati 1995; Rispail &

Leger 1998) and is currently classified in 31 genera and 35 subgenera (Marcondes 2007).

Phlebotominae are terrestrial breeders with larvae feeding on damp organic matter (Azar &

Nel 2003). The larvae are difficult to find in the field, but have been studied in laboratory cultures.

Adults of both sexes feed on nectar and honeydew; females additionally feed on vertebrate blood (Azar & Nel 2003).

Synapomorphies of the Phlebotominae include a complete reduction of A₁; the partition of tergite 9 into two long lateral lobes in the male; gonostylus pointed towards T9 and carrying characteristic thorns; and sperm pump without connection to aedeagus (Hennig 1972). Additional characteristics include the slender general appearance, the functional mandibles and maxillae of the female, the five-segmented palp and the elongated proboscis (Williams 1993).

The fossil record of Phlebotominae dates back to the early Cretaceous (e.g. Hennig 1972;

Azar et al. 1999), and several extant genera are known from Mexican, Dominican and Baltic ambers (Hennig 1972; Poinar 2008). Distributional data, however, point to the phlebotomines as an older group even than this (Ilango 2010). The oldest documented association with parasitic protozoans is from Burmese amber, believed to be from the late Cretaceous (Poinar 2004).

Psychodinae Newman, 1834

The Psychodinae are a highly derived group containing the majority of Psychodid species diversity.

About 2000 species have been described in about 100 genera distributed all over the world (Wagner et al. 2008). No satisfactory system of classification has yet been developed, and different authors often follow widely different generic and tribal concepts (e.g. Vaillant 1971; Duckhouse 1987;

Ježek & van Harten 2005).

The ecology of Psychodinae was summarised by Vaillant (1971), who classified them into 11 functional groups based on habitats. Most species are freshwater detritivores, but there are also species found in fungal fruit bodies, leaf litter and compost, decaying wood, dead snails and vertebrate faeces. A few species are opportunistic myiasis agents, but confirmed cases of this are rare (Smith & Thomas 1979; Taylan-Ozkan et al. 2004; Tu et al. 2007).

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The Psychodinae exhibit a number of synapomorphic characters including a well developed eyebridge (secondarily lost in some genera); reduced mouthparts; both basal cells of wings shortened; a triangular neala at wing bases; cerci of males modified into cercopods carrying retinaculae; gonopods directed at cercopods; tergite 9 fused with sternite 9 to form a ring; sessile spermathecae and elongated cerci present in the females; the larvae with 1-segmented antennae and breathing tube and pupae with basal part of prothoracal horn moveable (Hennig 1972).

Whereas the monophyly of the subfamily is well established, its relationships with other subfamilies remains unclear. Hennig (1972) and Azar et al. (1999), based on morphology of mainly fossil species, suggested a clade Trichomyiinae+Sycoracinae+Horaiellinae to be the sister group of the Psychodinae. On the other hand, Curler (2009) found the sister group to be Bruchomyiinae based on morphology of all life stages, and either Trichomyiinae or Trichomyiinae+Horaiellinae based on molecular phylogenies.

Fossil Psychodinae are found in many amber deposits, but remain poorly studied (Evenhuis 1994). A molecular clock analysis based on European species suggested the major radiations in the group to have taken place around 85 million years ago (Espindola 2010). There are, however several reasons to doubt this result as the mitochondrial markers used were putatively saturated and because only European species were included (Espindola 2010). An older age is also suggested by Gondwanan distributions in e.g. the Maruinini (Duckhouse 1990).

Sycoracinae Jung, 1954

The Sycoracinae is a species-poor group, consisting of only 36 described extant species (Santos et al. 2009a; Bravo et al. 2010). Although sometimes considered monogeneric (Ježek 1999), most authors recognise 3 genera in the subfamily (Duckhouse 1972; Santos et al. 2009a). The group has a near cosmopolitan distribution, lacking only in North America.

Females of many species have been found to be haematophagous on amphibians, and the European Sycorax silacea Haliday in Curtis, 1839 is a vector for microfilarial worms (Desportes 1941; Bravo & Salazar-Valenzuela 2009). Known immature stages are small and aselliform and are found in aquatic mosses or leaf litter (Duckhouse 1972; Wagner 1997).

Proposed synapomorphic characters of Sycoracines include a shortened CuA₂; sternites of the eight abdominal segment of the female present as a narrow ring separated from S9 and hypopygium of male not inverted relative to body axis (Duckhouse 1972). Other characteristic features include developed mandibles in females; R with four branches; basal cell elongated; palp with four segments and gonostylus with a terminal spine (Duckhouse 1972). Most of these non- unique characters are plesiomorphic in the Psychodidae, except the 4-branched R which may or may not be indicative of a relationship with Trichomyiinae. This relationship was hypothesised

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by Hennig (1972), and was also recovered in an 18S molecular study by Curler (2009). Other molecular analyses have, however, failed to replicate this result (Curler 2009).

The oldest Sycoracine fossils date back to the Cretaceous, and it is generally believed that the group’s current diversity is much lower than it was in the past (Azar et al. 2007).

Trichomyiinae Tonnoir, 1922

The Trichomyiinae form an ancient and heterogenous group consisting of around 100 described extant species (Bravo 1999). It is cosmopolitan in distribution, with species being found on all continents except Antarctica. The only extant genus is Trichomyia Haliday in Curtis, 1839; which has not yet been satisfactorily subdivided. Tentative classifications divide it into a «group A» and a

«group B» (Duckhouse 1965) and/or into six subgenera (Bravo 2001).

The larvae of several Trichomyiinae have been found to be xylophagous (Keilin 1914;

Duckhouse 1978), and it has been speculated that most or all species live this way (Wagner 1982).

The adults are often rare in collections, but this is due to inappropriate collecting methods more than actual rarity in nature (Duckhouse 1978). Males of many species are attracted to light and are best collected using light traps (Duckhouse 1978).

Apomorphic characters characterising Trichomyiinae include annulated spermathecal ducts with cup-like sceroterisations connected to the spermathecae and the larva’s adaptations to terrestrial, xylophagous feeding (Keilin & Tate 1937; Duckhouse 1980). Other characteristics include reduced mouthparts; palps with three or four segments; the inner part of the first or second palp segment with a group of sensillae; R with four branches; CuA₂ long and cerci similar in males and females (Duckhouse 1972; Hennig 1972; Wagner 1982).

About 20 fossil species of Trichomyiinae are known, belonging to the genera Trichomyia, Eatonisca Meunier, 1905 and Eotrichomyia Nel, Menier & De Ploëg, 2002 (Lak et al. 2008).

The oldest Trichomyia species known is from Cretaceous French amber, but the Gondwanan and Transantarctic distribution of some groups are suggestive of an even older age (Duckhouse 1972, 1980).

Horaiellinae Enderlein, 1936

The subfamily Horaiellinae was erected for the single genus Horaiella Tonnoir, 1933, and is known from four obscure species from India, China and Thailand (Curler et al. 2006). Like the Sycoracinae and the Trichomyiinae it has R with four branches, but whereas R₄ and R₅ are fused in these subfamilies, Horaiella has a fusion between R₂ and R₃ (Hennig 1972). As in the Sycoracinae and the Phlebotominae, the female has functional mouthparts. This is, however, most likely a

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plesiomorphic condition.

Larval Horaiellinae live in fast-flowing streams and in the splash zones of waterfalls. For attachment, they have large ventral suckers similar to those of the Psychodine genera Maruina Müller, 1895, Neotelmatoscopus Tonnoir, 1933 and Neomaruina Vaillant, 1963. However this similarity is surely superficial (Duckhouse 1985).

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Ježek, J. (1999) Comments on the correct grammatic gender of Sycorax Curt. and Philosepedon Eat. (Diptera: Psychodidae) with world catalogue. Dipterologica bohemoslavica, 9, 83-87.

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Ježek, J. & Hájek, J. (2007) Psychodidae (Diptera) of the Orlické hory Protected Landscape Area and its environs with descriptions of two new species from the Czech Republic. Acta Entomologica Musei Nationalis Pragae, 47, 237-285.

Ježek, J. & van Harten, A. (2005) Further new taxa and little-known species of non-biting moth flies (Diptera, Psychodidae) from Yemen. Acta Entomologica Musei Nationalis Pragae, 45, 199-220.

Keilin, D. (1914) Sur la biologie d’un Psychodide a Larve xylophage Trichomyia urbica CURTIS (Diptère). Comptes Rendus Hebdomaire des Seances et Memoires de la Societe de

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Leite, A. C. R., Williams, P. & Dossantos, M. C. (1991) The Pupa of Lutzomyia longipalpis (Diptera, Psychodidae - Phlebotominae). Parassitologia, 33, Suppl. Dec. 1991, 477-484.

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Catalogue of Afrotropical Psychodidae

Chapter 2

Catalogue and bibliography of Afrotropical Psychodidae:

Bruchomyiinae, Psychodinae, Sycoracinae and Trichomyiinae

INTRODUCTION

The first major contribution to non-phlebotomine Afrotropical Psychodidae were the studies of Tonnoir (1920, 1922, 1939a, 1939b), in which 34 Afrotropical moth fly species were described mostly from East Africa. Another important pioneer of the continent’s Psychodidae fauna was Satchell (1955), whose comprehensive key remains useful today despite its outdated nomenclature.

To date, the most important studies of the African Psychodidae fauna were conducted by Duckhouse (1975, 1978, 1980, 1985a, 1987), based mostly on collections by G. H. Satchell and B. Stuckenberg housed in the Natal Museum in Pietermaritzburg, South Africa. In addition to describing a total of 40 species, he also described and revised several genera, catalogued the fauna (Duckhouse &

Lewis 1980) and shed light on several systematic and biogeographic questions.

Other studies on the taxonomy of Afrotropical Psychodidae include Eaton (1913), Edwards (1929), Freeman (1949), Quate (1957), Stuckenberg (1962, 1978), Hogue (1970), Salamanna (1980), Wagner (1979a, 1983, 1989), Ježek (2004), Ježek & van Harten (1996, 2002, 2005, 2009) and Wagner & Andersen (2007). A total of 173 non-phlebotomine Psychodidae species are currently known from the region, however this is likely only a small fraction of the total fauna.

Africa’s moth fly fauna remains poorly studied and there is still much to be learned even from small collections (Ježek & van Harten 2005; Wagner & Andersen 2007).

The first check list of Afrotropical Psychodidae was presented in Tonnoir (1939) without bibliographic references for the individual species. Another early catalogue was prepared by Rapp

& Cooper (1945), but today this work only has historical value as its nomenclature is outdated and inconsequent. In Duckhouse & Lewis’ (1980) treatment of the family in the catalogue of Afrotropical Diptera, some, but not all of the nomenclatural errors were corrected. Since then, many new species and genera have been recorded from the Afrotropics and an updated catalogue will prove useful for future taxonomic endeavors in the region.

Systematic and geographic scope

The present catalogue covers the Afrotropical species of Psychodidae: Bruchomyiinae,

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Psychodinae, Sycoracinae and Trichomyiinae. Systematics of Psychodidae are generally stable on the subfamily level; however conflicting classifications exist within several of the subfamilies. This catalogue follows Duckhouse’s (1987) provisional tribe-level systematics of Psychodinae and is in most cases in accordance with Duckhouse & Lewis (1980) on generic nomenclature. It deviates mainly within the Mormiini and Paramormiini; the systematics of which have been the focus of much recent work by Ježek (e.g. 1983, 1984a, 1989, 1990, 1994, 2004). Some new nomenclatural acts are proposed; these are discussed in further detail below. Subfamilial and tribal autorship is according to Sabrosky (1999).

The geographic scope of this work is the Afrotropical region as defined in Crosskey (1980), modified according to the forthcoming Manual of Afrotropical Diptera (Kirk-Spriggs & Mostovski, in prep.). Accordingly, the Afrotropical region is recognized as Africa South of the Sahara (starting for convenience with the Northern boundaries of Mauritania, Mali, Niger, Chad and Sudan), the Cape Verde Islands, Madagascar, the islands of the Southern Indian Ocean and the Southernmost part of the Arabian peninsula including Yemen, Oman and the United Arab Emirates.

Format

The catalogue lists Afrotropical subfamilies, tribes, genera, subgenera and species of Afrotropical Psychodidae exclusive of Phlebotominae. All supraspecific taxa are written in capitals. Valid names are listed in bold, with synonyms and unavailable names in regular typeface. Each entry contains the following bibliographic records: Author, year, title of publication, issue and first page number of taxon description. Type species with type fixation criteria are given for genera. Species are listed with type localities and distributional information based on the published literature. For species described in genera other than those in which they currently placed, the original genus is given in italics.

Type localities are mostly quoted directly from the original descriptions. For a few species described from other zoogeographical regions, the type localities are quoted from previous Psychodidae catalogues, notably Wagner (1990). In cases where names of type localities are ambiguous or outdated the modern names of the respective countries are given in square brackets.

The Democratic Republic of Congo is abbreviated D.R. Congo and the United Arab Emirates are abbreviated UAE.

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Catalogue of Afrotropical Psychodidae

CATALOGUE

Subfamily BRUCHOMYIINAE ALEXANDER, 1921

BRUCHOMYIINAE ALEXANDER, 1921 (as subfamily of Tanyderidae): Annals of the Entomological Society of America 13 (1920): 402.

NEMOPALPINAE EDWARDS, 1921: Annals and Magazine of Natural History 7: 439 Genus EUTONNOIRIA ALEXANDER, 1940

EUTONNOIRIA ALEXANDER, 1940: Revista de Entomologia 11: 794. Type species: Bruchomyia edwardsi Tonnoir, 1939b (original designation)

edwardsi (TONNOIR, 1939b): Ruwenzori Expedition 1: 38 (Bruchomyia). Type locality: [Uganda]

«RUWENZORI: Mobuku valley, 7300 ft.» - Distr.: Uganda.

Genus NEMAPALPUS MACQUART, 1838

NEMAPALPUS MACQUART, 1838: Mémoires de la Société royale des sciences, de l’agriculture et des arts de Lille 1: 85. Type species: Nemapalpus flavus Macquart, 1838 (monotypy)

NEMOPALPUS MACQUART, 1839: Histoire Naturelle des Iles Canaries 2: 102. Variant spelling.

NYGMATODES LOEW, 1845: Dipterologische Beiträge 1: 9. Unavailable name.

PALAEOSYCORAX MEUNIER, 1905: Miscellana Entomologica 13: 50. Type species: Palaeosycorax tertiariae Meunier, 1905 (monotypy)

capensis EDWARDS, 1929: Annales and Magazine of Natural History 3: 422. Type locality: «S.

AFRICA: Port St. John, Pondoland» - Distr.: South Africa

concolor STUCKENBERG, 1962: Annals of the Natal Museum 15: 211. Type locality: [South Africa]

«Port St. Johns, Pondoland, Eastern Cape Province» - Distr.: South Africa

davidsoni STUCKENBERG, 1978: Annals of the Natal Museum 23: 372. Type locality: [Namibia]

«SOUTH WEST AFRICA. Windhoek District, Daan Viljoen Park, approximate lat. 22°30’S, long.

16°55’E, altitude c. 1 900 m.» - Distr.: Namibia

ledgeri STUCKENBERG, 1978: Annals of the Natal Museum 23: 369. Type locality: [Namibia] «SOUTH WEST AFRICA, Bethanie Dist., Oberndorf farm, approx. lat. 26°22’S, long. 17°09’E» - Distr.:

Namibia

transvaalensis STUCKENBERG, 1962: Annals of the Natal Museum 15: 211. Type locality: [South Africa] «Mariepskop, Pilgrims Rest District, Eastern Transvaal» - Distr.: South Africa

Subfamily PSYCHODINAE NEWMAN, 1834 PSYCHODITES NEWMAN, 1834: Entomological Magazine 2: 379.

Tribus MARUININI ENDERLEIN, 1937

MARUININI ENDERLEIN, 1937: Deutsche Entomologische Zeitsschrift 1936: 109 ARISEMINI VAILLANT, 1982a: Nouvelle Revue d’Entomologie 12: 190

SETOMIMINI VAILLANT, 1982a: Nouvelle Revue d’Entomologie 12: 191

Genus NEOARISEMUS BOTOSANEANU & VAILLANT, 1970

NEOARISEMUS BOTOSANEANU & VAILLANT, 1970: Travaux du Laboratoire d’Hydrobiologie et de Pisciculture de l’Universite de Grenoble 61: 178. Type species: Psychoda nigra Banks, 1894 (original designation)

advenus DUCKHOUSE, 1978: Annals of the Natal Museum 23: 321. Type locality: «SOUTH AFRICA, Cape Province, Grahamstown, Chalmers Waterfall» Distr.: South Africa

anarticulatus DUCKHOUSE, 1978: Annals of the Natal Museum 23: 315. Type locality: «SOUTH AFRICA, Cape Province, near Knysna, Bracken Hill stream». Distr.: South Africa

(30)

angularis DUCKHOUSE, 1987: Annals of the Natal Museum 28: 237. Type locality: «SOUTH AFRICA, Cape Province, Garden of Eden, forest, 342/3AA.» Distr.: South Africa

brevicornis DUCKHOUSE, 1978: Annals of the Natal Museum 23: 329. Type locality: «SOUTH AFRICA, Cape Province, Stutterheim, Kologha Forest.» Distr.: South Africa

brunneus DUCKHOUSE, 1987: Annals of the Natal Museum 28: 242. Type locality: «TANZANIA, 6,5 miles S of Morogoro, 490 m, wet forest». Distr.: Tanzania

collarti (SATCHELL, 1955b): Revue de zoologie et de botanique africaines 51: 350. (Telmatoscopus).

Type locality: [D.R. Congo] «Belgian Congo : Stanleyville». Distr.: D.R. Congo

deviatus (TONNOIR, 1939b): Ruwenzori Expedition 1:46 (Psychoda). Type locality: [Uganda]

«RUWENZORI: Bwamba Pass (West side) 5500-7500 ft.». Distr.: Uganda.

elongatus DUCKHOUSE, 1978: Annals of the Natal Museum 23: 317. Type locality: «SOUTH AFRICA, Natal Province, Drakensberg Mts. (2929 Ad), Giant’s Castle Reserve, 1 768 m». Distr.: South Africa

impeditus DUCKHOUSE, 1978: Annals of the Natal Museum 23: 332. Type locality: «SOUTH AFRICA, N.E. Transvaal, Tzaneen, Magoeba’s Kloof.» Distr.: South Africa, Zimbabwe

obtusistylus DUCKHOUSE, 1978: Annals of the Natal Museum 23: 320. Type locality: «SOUTH AFRICA, N.E. Transvaal, Tzaneen, Magoeba’s Kloof.» Distr.: South Africa

pectinatus (TONNOIR, 1939b): Ruwenzori Expedition 1:66 (Telmatoscopus). Type locality: [Uganda]

«RUWENZORI: Namwamba Valley, 10,200 ft..». Distr.: Uganda.

plesius DUCKHOUSE, 1978: Annals of the Natal Museum 23: 313. Type locality: «SOUTH AFRICA, Cape Province, Ashton, Klaas Vooges Kloof». Distr.: South Africa

pristinus DUCKHOUSE, 1987: Annals of the Natal Museum 28: 239. Type locality: «SOUTH AFRICA, Natal, Pietermaritzburg, Town Bush». Distr.: South Africa

prodigiosus DUCKHOUSE, 1978: Annals of the Natal Museum 23: 310. Type locality: «SOUTH AFRICA, near Cape Town, Kalk Bay stream». Distr.: South Africa

propensus (JUNG, 1956): Deutsche Entomologische Zeitschrift, N.F. 3: 185 (Telmatoscopus, as new name for Telmatoscopus fuscus Tonnoir, 1939 nec (Macquart, 1826)). Type locality: [Uganda]

«Namwamba Valley, 6500 ft.». Distr.: Uganda.

fuscus (TONNOIR, 1939b) nec Macquart, 1826: Ruwenzori Expedition 1:65 (Telmatoscopus). Type locality: [Uganda] «Namwamba Valley, 6500 ft.». Distr.: Uganda.

satchelli DUCKHOUSE, 1978: Annals of the Natal Museum 23: 324. Type locality: «SOUTH AFRICA, Cape Town, Kirstenbosch, Skeleton Stream, upper reaches.» Distr.: South Africa

tapetipennis DUCKHOUSE, 1978: Annals of the Natal Museum 23: 326. Type locality: «SOUTH AFRICA, Cape Town, Kirstenbosch, Skeleton Stream, upper reaches.» Distr.: South Africa youngi DUCKHOUSE, 1987: Annals of the Natal Museum 28: 240. Type locality: «TANZANIA, 6,5

miles S of Morogoro, 490 m, wet forest». Distr.: Tanzania

Genus SETOMIMA ENDERLEIN, 1937

SETOMIMA ENDERLEIN, 1937: Deutsche Entomologische Zeitschrift 4: 100.Type species: Setomima lithocolleta Enderlein, 1937 (original designation)

PARABRUNETTIA VAILLANT, 1975: Die Fliegen Der Palearktischen Region 310: 165 (as subgenus of Brunettia Annandale, 1910). Type species: Psychoda nitida Banks, 1901 (original designation)

Subgenus OPHRYOSETOMIMA DUCKHOUSE, 1987

OPHRYOSETOMIMA DUCKHOUSE, 1987: Annals of the Natal Museum 28: 255 (as subgenus of Setomima Enderlein, 1937.Type species: Setomima spinifera Duckhouse, 1978 (original designation)

brachiata DUCKHOUSE, 1987: Annals of the Natal Museum 28: 260. Type locality: «KENYA, Rift Valley, Kampi-Ya-Samaki, Lake Baringo, 00°37´N:36°02´E, 980 m.». Distr.: Kenya, South Africa.

longispinosa DUCKHOUSE, 1987: Annals of the Natal Museum 28: 256. Type locality: «SOUTH AFRICA, Natal, Eshowe District, Dlinza Forest, 450». Distr.: South Africa.

Biodiversity studies in Afrotropical moth flies (Diptera: Psychodidae)